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Generic Adjustments in Neotropical Malpighiaceae

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Generic Adjustments in Neotropical Malpighiaceae 2007 Contr. Univ. MichiganANDERSON Herb. 25: 137–166. & DAVIS: 2007. NEOTROPICAL MALPIGHIACEAE 137 GENERIC ADJUSTMENTS IN NEOTROPICAL MALPIGHIACEAE William R. Anderson University of Michigan Herbarium 3600 Varsity Drive, Ann Arbor, Michigan 48108-2287 Charles C. Davis Department of Organismic and Evolutionary Biology Harvard University Herbaria 22 Divinity Avenue Cambridge, Massachusetts 02138-2094 ABSTRACT. The genera Bronwenia W. R. Anderson & C. Davis and Calcicola W. R. Anderson & C. Davis are proposed to accommodate species segregated from Banisteriopsis and Mascagnia, respectively, and Cottsia Dubard & Dop is resurrected for the North American species formerly assigned to Janusia. All three genera are described, their morphology and relationships are discussed, and keys to the spe- cies are provided, with nomenclature for each species. For Calcicola and Cottsia distribution maps and descriptions of species are provided. The genus Clonodia is placed in synonymy under Heteropterys, with discussion of relationships, a key to clonodioid species, and nomenclature and a diagnostic description for each species. One new species is described [Bronwenia peckoltii W. R. Anderson & C. Davis] and 19 new combinations are proposed [Bronwenia acapulcensis (Rose) W. R. Anderson & C. Davis, B. acapulcensis var. llanensis (B. Gates) W. R. Anderson & C. Davis, B. brevipedicellata (B. Gates) W. R. Anderson & C. Davis, B. cinerascens (Benth.) W. R. Anderson & C. Davis, B. cornifolia (H. B. K.) W. R. Anderson & C. Davis, B. cornifolia var. maracaybensis (Adr. Juss.) W. R. Anderson & C. Davis, B. cornifolia var. standleyi (B. Gates) W. R. Anderson & C. Davis, B. ferruginea (Cav.) W. R. Anderson & C. Davis, B. longipilifera (B. Gates) W. R. Anderson & C. Davis, B. mathiasiae (W. R. Anderson) W. R. Anderson & C. Davis, B. megaptera (B. Gates) W. R. Anderson & C. Davis, B. wurdackii (B. Gates) W. R. Anderson & C. Davis, Calcicola parvifolia (Adr. Juss.) W. R. Anderson & C. Davis, C. sericea (Nied.) W. R. Anderson & C. Davis, Cottsia californica (Benth.) W. R. Anderson & C. Davis, C. gracilis (A. Gray) W. R. Anderson & C. Davis, C. linearis (Wiggins) W. R. Anderson & C. Davis, Heteropterys complicata (H. B. K.) W. R. Anderson & C. Davis, and H. ovata (Nied.) W. R. Anderson & C. Davis]. Ten species are illustrated. INTRODUCTION Phylogenetic studies of Malpighiaceae published in recent years (Cameron et al. 2001; Davis et al. 2001, 2002), and our recently acquired unpublished data, have revealed a number of genera that need to be divided or combined with other genera to produce a more nearly monophyletic classification. The purpose of this paper is to deal with four of those problems. Although we give many synonyms below, we are omitting some of the less important infraspecific names; complete synonymy can be obtained by consulting the nomenclatural database on the Malpighiaceae website at http://herbarium.lsa. umich.edu/malpigh. BANISTERIOPSIS AND BRONWENIA In her monograph of Banisteriopsis C. B. Rob. and Diplopterys Adr. Juss., Gates (1982) recognized three subgenera in Banisteriopsis and considered Diplopterys a separate genus resembling Banisteriopsis subg. Pleiopterys (Nied.) B. Gates. Anderson 137 138 CONTR. UNIVERSITY OF MICHIGAN HERBARIUM VOLUME 25 and Davis (2006) reported that Banisteriopsis sensu Gates is not monophyletic, and that Diplopterys is nested within Banisteriopsis subg. Pleiopterys. We resolved that problem by transferring all the species of Banisteriopsis subg. Pleiopterys to Diplo- pterys. The remaining two subgenera of Gates, however, require further taxonomic elucidation. Some clarification of the subgeneric nomenclature is in order. Gates assumed that the earliest designation of a lectotype for Banisteriopsis was that of Cuatrecasas (1958, p. 486), who treated B. cornifolia (H. B. K.) C. B. Rob. as the “type” [lectotype]. On that basis Gates used the name subg. Banisteriopsis for the group of eight spe- cies containing B. cornifolia. The other two, much larger, subgenera she called subg. Hemiramma (Griseb.) B. Gates and subg. Pleiopterys. We now know, however, that there was an earlier designation of a lectotype for Banisteriopsis by Morton (1931), B. argentea (H. B. K.) C. B. Rob. [B. muricata (Cav.) Cuatrec.]. Banisteriopsis muricata belongs to the very large subgenus Hemiramma, so that has to become subg. Baniste- riopsis and the generic name Banisteriopsis will remain (fortunately) with that largest group of species. The other, small subgenus that Gates called subg. Banisteriopsis is the subject of these notes. Our preliminary phylogenetic results (C. Davis & W. R. Anderson, unpubl.) indicate that the species assigned by Gates to her subg. Ban- isteriopsis do not form a clade with the greatly expanded subg. Banisteriopsis, and thus should not be retained in Banisteriopsis sens. str. (see Fig. 1, where the species in question are in boldface under the new name Bronwenia). We here segregate the eight species treated by Gates as Banisteriopsis subg. Banisteriopsis as a new genus, and include two additional ones not mentioned in her monograph. Bronwenia W. R. Anderson & C. Davis, gen. nov.—TYPE: Bronwenia ferruginea (Cav.) W. R. Anderson & C. Davis. Banisteria sect. Monoctenia Nied., Ind. Lect. Lyc. Reg. Hos. Brunsberg. p. hiem. 1900–1901: 4. 1900.—LECTOTYPE, designated by Gates, 1982, p. 41: Banisteria maracaybensis Adr. Juss. [Bronwenia cornifolia var. maracaybensis (Adr. Juss.) W. R. Anderson & C. Davis]. Lianae lignosae vel frutices; petiolus eglandulosus vel prope apicem biglandu- losus; lamina eglandulosa vel margine glandulis instructa; stipulae interpetiolares, distinctae; inflorescentia paniculata vel dichasialis, floribus in pseudoracemis 6–45- floris portatis; pedicelli sessiles vel brevipedunculati; glandulae calycinae pro parte maxima infra sepala affixae; petala lutea, plerumque glabra; antherarum connectivum loculos non superans; stigmata terminalia; samarae nux in lateribus laevis vel 1 crista vel alula instructa, ala dorsali elongata, adaxialiter incrassata; carpophorum nullum vel usque ad 1.5 mm longum, 1–2 mm latum. Woody vines or shrubs, when shrubby the branches often twining. Leaves decus- sate; lamina with glands none or usually marginal (but see B. peckoltii), sessile, the distal glands (if present) minute, the 2 most proximal glands usually enlarged; peti- ole eglandular or biglandular at or somewhat below apex; stipules usually present, interpetiolar, distinct, triangular, minute (0.2 mm long) or small (up to 1.5 mm long). Inflorescence axillary or terminal, paniculate or dichasial, the flowers borne in short, usually dense pseudoracemes of 6–45; bracts and bracteoles eglandular, persistent; pedicels sessile or short-pedunculate. Flowers bilaterally symmetrical in calyx and corolla, and sometimes in androecium. Sepals leaving petals exposed during enlarge- ment of bud, the lateral 4 bearing large paired abaxial glands and the anterior eglan- dular, the glands attached for most of their length below free part of sepal. Petals 2007 ANDERSON & DAVIS: NEOTROPICAL MALPIGHIACEAE 139 FIG. 1. Phylogenetic tree showing position of Bronwenia in the Stigmaphyllon clade of Malpighia- ceae. Bootstrap values are given above branches. The reduced strict consensus tree shown here is extracted from a larger phylogenetic analysis of Malpighiaceae using combined DNA sequences from plastid matK, ndhF, and rbcL, nuclear PHYC, and morphology; all genes were sequenced for all taxa (C. Davis & W. R. Anderson, unpubl.). Vouchers for species of Bronwenia sequenced: Bronwenia cornifolia: Clark 1, MICH; Bronwenia mathiasiae: Nelson et al. 518, MICH. yellow, glabrous (very rarely sericeous on abaxial midrib), the posterior petal dif- ferent in size, shape, and stance from the lateral 4, and its margin often more deeply divided with the divisions (especially proximally) ± glandular-thickened. Receptacle glabrous on both sides of filaments. Stamens 10, all fertile; anthers glabrous or pilose, the connective not or hardly exceeding locules at apex. Gynoecium 3-carpellate, the carpels 1 anterior and 2 posterior, all fertile; styles 3, distinct, mostly ± alike; stigmas terminal, truncate or capitate. Fruit dry, breaking apart at maturity into 3 samaras (or fewer by abortion) separating from a low pyramidal torus; dorsal wing of samara well developed, elongated, thickened on the adaxial edge with the veins bending toward the thinner abaxial edge, a shallow triangular or rounded appendage usually present on adaxial edge at base; nut of samara smooth-sided or bearing a single ridge or winglet on each side parallel to areole; locule of nut glabrous within; carpophore absent or present but short (up to 1.5 mm long), broad (1–2 mm wide), and non-func- tional. Chromosome number: n = 10 (in B. acapulcensis var. llanensis [Gates 1982]). Bronwenia is notable for its leaf glands usually marginal (if present), flowers borne in short dense pseudoracemes, calyx glands attached below the free part of the sepals, yellow, usually glabrous petals, and samaras with the sides of the nut smooth or bearing a single ridge or winglet parallel to the areole and the carpophore absent or short, broad, and apparently non-functional. Of the ten species recognized here, the eight that were treated in the same subgenus by Gates are morphologically coher- ent and readily recognized by their small petals. For descriptions of those species and their varieties, see Gates (1982). The two additional species of Bronwenia (B. mathiasiae and B. peckoltii) have
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